Jurnal Natur Indonesia 13(3), Juni 2011: 226-231 226ISSN 1410-9379,Jurnal KeputusanNatur Indonesia Akreditasi13(3): No 226-23165a/DIKTI/Kep./2008 Siregar, et al. The Genus Licuala Wurmb () in Java

Etti Sartina Siregar1*) dan Sri Sudarmiyati Tjitrosoedirdjo2)

1)Departement of Biology, Faculty of Mathematics and Natural Sciences, University of Sumatra Utara, Medan 20155 2)Departement of Biology, Faculty of Mathematics and Natural Sciences, Bogor Agriculture Institute, Bogor 16680

Diterima 07-12-2010 Disetujui 07-03-2011

ABSTRACT The comprehensive revision of the genus has been done in the Malay Peninsula, New Guinea, Sumatra and Kalimantan. In Java, the revision of Licuala has never been recently completed. This study was based on morphological characters of specimens which are preserved in Herbarium Bogoriense. The several living collections cultivated in Bogor Botanical Garden were studied. The aim of this study is to ascertain the correct names and improve the species delimitation of Licuala in Java. The result shows that there are three species of Licuala in Java. They are L. gracilis, L. pumila and L. spinosa. Two previously known species have been placed into synonymies; L. flabellum to L. gracilis, and L. spectabilis to L. spinosa. Licuala gracilis is endemic for west Java.

Keywords: Arecaceae, Java, Licuala

INTRODUCTION Myanmar, Bangladesh, and India (north-eastern part and Licuala is a genus belonging to the family the Andaman Islands). The centre of species diversity Arecaceae (a conserved alternative name for the consider is at the two main areas, one within the Sunda Palmae), subfamily , tribe Corypheae and Shelf in Malaya and Borneo and the other at the Sahul sub tribe Livistoninae (Uhl & Dransfield, 1987). It is a Shelf on the island of New Guinea (Saw, 1997; Saw small to medium sized tree palm, rarely arborescent et al., 2003). and easily recognized by the wedge-shaped fronds with The name Licuala is derived from the Makassar segments that split to the hastula. Naturally, all spe- (Celebes) name ‘leko wala’ (Backer, 1936). It was first cies of Licuala are understorey palms. In general, the described by Rhumpius (1741), in the Herbarium genus does not survive under open conditions and com- Amboinense as Licuala arbor. As this name was pre- monly found in the lowland forest. However, some spe- Linnean, so the name L. spinosa as was described by cies are also found in the other forest types, such as L. Wurmb (1780), was accepted by International Code of spinosa in open moist place and L. glabra in the hill Botanical Nomenclature or ICBN (Greuter et al., 2000). stations. Traditionally, some species of this genus such Thus, L. spinosa is regarded as the genus type of as L. acutifida, L. grandis, L. paludosa and L. spinosa, Licuala. are used as ornaments, decoration, roofing, food-wrap- The genus contains wide morphological variations pers, walking sticks, binding, making hats and eaten and these cause a serious problem to the researcher in as vegetable (Burkill, 1935; Whitmore, 1973; Dransfield, the process of revising. These morphological variations 1976; Saw, 1997). were easily observed in the field rather than through The genus consists of 141 species, occuring from herbarium specimens. Thus, taxonomic study is based Vanuatu, the Solomon Islands, New Guinea, Australia, mainly on the data gained from fieldworks, such as in- Indonesia, the Philippines, Malaysia, Thailand, florescences and flowers, which are not always found Cambodia, Laos, Vietnam, China (southern part), in the herbarium specimens. The last comprehensive revision of the genus in the Malay Peninsula has been done by Saw (1997),

*)Telp: +6281264852784 Email: [email protected] The genus Licuala Wurmb (Arecaceae) in Java 227 in New Guinea by Barford (2000), in Sumatera by Ardan species concept is adopted in this study, following (2000) and in Kalimantan by Batoro (2001). In Java, the Dransfield (1999). The terminologies followed Stearn revision of Licuala has never been recently completed. (1966), Uhl and Dransfield (1987), Dransfield and Beentje According to Blume (1836), there are two species of (1996). The basic description follows Uhl and Dransfield Licuala in Java, L. gracilis Bl. and L. pumila Bl. Miquel (1987), Saw (1997), Barfod and Saw (2002). (1851), reported 3 species, L. pumila Bl., L. spinosa Wurmb and L. spectabilis Miq. Koorders (1913), re- RESULTS AND DISCUSSIONS ported two species, L. pumila Bl. and L. spinosa Wurmb. Vegetative Organs. Licuala is rarely treelike, Backer & Bakhuizen van den Brink Jr. (1968), recog- small to medium sized, acaulescent to shrubby, soli- nized only two species of Licuala in Java, L. spinosa tary or clustered. Licuala pumila is solitary, whereas L. Wurmb and L. pumila Bl. In Bogor Botanical Garden, gracilis and L. spinosa are clustered. there are two clusters of living collection, which are be- is the tallest within the genus and almost treelike. lieved (to be) of Ujung Kulon original and identified as L. Licuala pumila is almost acaulescent. flabellum (Anonymous, 1985). Interestingly, this spe- Petioles are usually with spines. The spines are cies was previously described by Martius (1838), that found almost at the whole length of the petiole in L. was found in the wild of Celebes, however no other col- spinosa, about half of the petiole in L. gracilis and L. lections were found. pumila. Fronds are orbicular in L. gracilis and L. pumila, Therefore, it is considered essential to proceed the but peltate-orbicular in L. spinosa. The base of most study of this genus in Java to ascertain the correct fronds is equal but unequal in L. gracilis. The leaves are names and improve the species delimitation. The study always split into many segment. Central segment is is based on the herbarium materials from Herbarium usually entire, except L. gracilis, which has bifid Bogoriense (BO) and fresh materials from Bogor Bo- central segment. The central segment is slightly larger tanical Garden. than the rest in L. gracilis or about equal in size with the lateral segment in L. spinosa and L. pumila. MATERIALS AND METHODS Generative Organs. The inflorescence pattern Specimens of Licuala which were collected from provide some good characters in distinguishing the Java by the previous collector, fresh materials from species (Figure 1). The inflorescence is erect to patent, Bogor Botanical Garden and some other areas in Java longer than petiole in L. spinosa or shorter than petiole were the major materials for examinations. Collecting in L. gracilis and L. pumila. The inflorescence varies in method followed Dransfield (1986). Morphological the number of branching orders, from one in L. pumila

A B C

Figure 1. Inflorescence patterns of Licuala in Java: A. L. gracilis (from Dransfield 1498); B. L. pumila (from JPM 7723); C. L. spinosa (from Koorders 25443, Dransfield 3525) 228 Jurnal Natur Indonesia 13(3): 226-231 Siregar, et al. to two in L. spinosa and L. gracilis. Peduncular bract is inflorescence appear on different individuals). Stem about present or sometimes absent in L. Spinosa, but it is 1-3 m tall, to c. 20 cm diameter, brown, nodes to completely absent in L. gracilis as well as in L. pumila. 1.5 cm distant. Leaves 10-14 in crown, sheaths to 30 Flowers are hairy or glabrous. The flowers of L. cm with red - brown fibres; petioles to 120 cm long, spinosa is hairy, but glabrous in L. gracilis and L. pumila. thornly below, 10-12 mm near base, 5-6 toward apex, The flowers are sessile and solitary in L. Pumila, green; spines along one third to more lenght of petiole, whereas in L. gracilis and L. spinosa are arranged in triangular to claws, patent to reflexed, largest near the solitary or group. base; fronds peltate-orbicular, unequal at the base, c. The shape of calyx varies from cylindrical to 30-45 cm long, c. 60-80 cm wide, segment c. 9-17 cyathiform in L. spinosa, campanulate in L. pumila and leaflets, the size is not same (wider toward central); tubular in L. gracilis. Corolla usually exceeding the lateral segment 2-5 costulate, 25-44 x 2.5-5 cm; calyx, tubular at the base, thick with lobe acute, central segment slightly larger than rest, bifid 1/5-1/3 except in L. gracilis which has blunt apex. Staminal frond lenght, c. 10-14 costulate, c. 30-45 x 7-10 cm. ring is present in all species. Ovary shape are varies, Inflorescences erect, shorter than leaves (very short), from turbinate in L. pumila and L. spinosa to cone shape c. 15-40 cm long, extending beyond crown (axilary), in L. gracilis. It is glabrous in all species under study. branching to two order, bears either entirely female or Fruit mostly globose with smooth surface. hermaphrodite flowers at a different individual; prophyll Immatured fruits are pink or cherry red in L. gracilis, tubular, c. 10-15 cm long, c. 1.5 cm wide, coriaceus whereas the other species have green immature fruits. and flattened at the base, closely sheathing, glabrous The seeds have the same shape as the fruits. The at the base and covered with brown hairs at the apex; surface is always smooth. peduncle c. 10-12 cm long, c. 2-3 cm wide, covered TAXONOMY with sparsely golden brown hairs, peduncular bract Key To The Species Of Licuala In Java lacking; rachis bracts only two, c. 10 cm long, c. 5 mm 1. Habit solitary .……...... ……….. 2. L. pumila wide; rachillae 8-28, unornamented, c. 8-18 cm long, c. Habit cluster ……...... ……………...2 2-3 mm wide, covered with sparsely golden brown hairs. 2. Inflorescence longer than petiole, outer surface of Flowers sessile, maturing not simultaneously, bud c. calyx and corolla hairy, young fruit green 4-5 x 2 mm; calyx tubular, c. 2.5-3 x 2-2.5 mm, base ...... 3. L. spinosa thickened, apex trilobed, acuminate, glabrous, Inflorescence shorter than petiole, outer surface of yellowish in colours; corolla tripartitus, triangular, c. calyx and corolla glabrous, young fruit pink or cherry 3.5-5 x 2 mm, base thickened, glabrous, apex blunt, red …………...... 1. L. gracilis green yellowish in colour; pistillate flowers solitary, 1. Licuala gracilis Blume.- Figure 2. ustering, spirally arranged, stays in anthesis for 8 to 10 days; gynodioecy (the hermaphrodite and female ovary cone shaped, glabrous, to 3.5 mm long and 1.4 mm wide; hermaphrodite flowers arranged in group of 2-4 flowers, densely arranged, cincinni 10-15 (c. 4 clusters per cm), stay in anthesis for 3 to 4 weeks; staminal ring membranous, c. 1mm long, filament subulate, c. 1 mm, base thickened; anther c. 1 mm long; ovary cone shaped, glabrous, c. 2 x 0.8 mm, consist of three uniovulate carpels, all three carpels in every flower develop into independent fruits (Figure 2). Fruit globose c. to 9 mm in diameter, glabrous, pink or cherry red in young and dark red or dark brown when Figure 2. L. gracilis (from Dransfield 1498);a-d; female flowers; mature (the colour variations occur in the cluster, not in a. Rachilla with some flower buds; b.open flower bud; c.ovary; d. carpel in cross section. e-i. hermaphprodite individual), solitary or develops an apocarpous fruit with flowers; e. rachilla with some flower buds; f-g. open three drupes (sometimes one flower bears three fruits); flower buds showing the stamens; h. ovary; i. carpel in cross section. j. The apocarpous fruit with three drupes; seed globose c. 6- mm in diameter, dark brown and k. one drupe of fruit; l. one drupe in section smooth. The genus Licuala Wurmb (Arecaceae) in Java 229

Distribution: Endemic to West Java. Habitat: pri- ment entire, 4-7 costulate, 33 x 35 cm. Inflorescences mary forest, common forest undergrowth component, shorter than leaves, erect to patent, 40-55 cm long with seashore to mountain about 150-750 m altitude. 2-3 partial inflorescences; branching to one order; one Notes: Based on the description of L. flabellum in to two opposite branching in proximal part, the upper Martius (1838), it has two similar characters with L. usually occupied by a single rachilla; peduncular bract gracilis, which are the bifid central segment and the lacking; peduncle tubular, flattened at the base, inflorescence pattern. This species was described by c. 10 cm long, c. 5 mm wide, glabrous, green in colour; Martius based on Reinwardt collections which was said rachis strongly flattened, glabrescent or sometime originated from Celebes. However, no collections have slightly puberulous; rachillae 4-5, c. 5-10 x 2 cm size, ever been made. So, there was a collection of this covered with sparsely golden brown hairs, bracts c. from Celebes. The type of L. flabellum is kept in Munich. 4-7 cm. Flowers solitary, irregularly, sessile, bud size 6 According to Dransfield and Moore (1982), the entire x 2.5-3 mm; calyx cylindrical to cyathiform, c. flowers have fallen from the type and in many cases 4 x 3 mm, very slightly three toothed, base thickened, Reinwardt had written the location for his specimens apex acuminate, glabrous, striate externally, green in incorrectly. Therefore, so they suggest that the type of colour; corolla coriaceus, triangular, tripartitus, c. L. flabellum has probably been collected from Ujung 5 x 3 mm, thick, glabrous, base thickened, apex acute, Kulon. Thus, it is regarded as synonym. striate externally, deeply sculptured inside, whitish in Specimen examined: JAVA: Ujung Kulon - Taman colour; staminal ring membranous, filament subulate, Jaya, Mc Donald & Afriastini 3378, fr., 18-5-1992 (BO!); anthers c. 0.5 mm long, blunt; ovary turbinate, glabrous, G. Payung, Kosterman 21846, fl+fr., 18-11-1960 (BO!, sculptured above, c. 2 mm long; style subulate, c. K, L); UNESCO 170, fl. ♂ + fr, s.d. (BO!); G. Kendeng, 1 mm long. Fruit sub globose, c. 9-10 x 8-9 mm, smooth, Kosterman 197, fl. ♀ , fl. ♂ + fr, 18-11-1960 (BO!), green in young and orange to red when mature. Seed Dransfield 1433, fl., 15-4-1971 (BO!, I, K, L, Sing), dark brown, sub globose c. 6-8 mm diameter, smooth. Dransfield 1498, fl. + fr., 21-4-1971 (BO!); Bogor Botanic Distribution: Sumatera and Java. Habitat: Ridge Garden - cultivated, Etti Siregar 06, fl. ♀ +fr and Etti top hill of Dipterocarp forest to lowland Dipterocarp for- Siregar 07, fl. ♂ + fr., 19-6-2003 (BO!). ests, 90 - 1000 m altitude. Specimen examined: JAVA: 2. Licuala pumila Blume. - Figure 3. Solitary, Lebak - Sunarari River, Backer 6362, fl., 1-1-1913 (BO!), acaulescent or with short stem no more than 1 m tall, Bakhuizen van den Brink 7946, fr., 1-4-1933 (BO!); about 1.5-2 cm in diameter, pale brown in colour. Leaves Bojong Manik- G. Liman, Koorders 40879â, fr., 14-7 - in crown 10-14; sheaths brownish; petioles up to 1.5 m 1912 (BO!); Jasinga: Dungus Iwul, Etti Siregar 03, fr., long, 5-7 mm wide near base, c. 2.2 mm wide toward 20-4-2003 (BO!); Steenis 11170, fr., 18-12-1938 (A, BO!, apex, greenish colour, with sparsely spines along less L, Sing), Dransfield 1506, fr., 15-5-1971 (BO!, I); G. half of petiole, little and turn, claws shape, dense near Halimun, JPM 7723, fl., th. 2003 (BO!); Cigelung, base of petiole, c. 3-4 mm long and 0.5-1 mm wide. Beümee A367, fl., 150, 23-11-1926 (BO!), Backer 10232, Fronds orbicular, apex shallow dentate, radiately parted fr., 24-11- 1913 (BO!); Janlappa, Meijer 2939, st., 20-8- 7-14 or more up to 18, c. 33 cm long, 41 cm wide, 1954 (A, BO!, K, L, Sing), Dransfield 991, fr., 11-10- lateral segment 2-4 costulate, 31 x 4.5 cm, central seg- 1970 (BO!); Leuliang, Bakhuizen van den Brink 7915,

Figure 3. L. pumila (from JPM 7723, Etti Siregar 03); a. Rachilla with some flower buds; b.flower bud; c. calyx and Figure 4. L. spinosa (from Koorders 25443, Dransfield 3525); corolla removed revealing the androecium; d. ovary; e. a. Rachilla with some fruits; b.flower bud; c.flower carpel in cross section. f. young fruit; g. young fruit in with calyx removed; d. calyx and corolla removed section revealing the androecium; e. ovary; f. young fruit 230 Jurnal Natur Indonesia 13(3): 226-231 Siregar, et al. st., 30-11-1932 (BO!); Sukabumi: Lengkong, Dransfield Java, Borneo. Habitat: Open, slightly swampy ground, 1062, fr., 25-11-1970 (BO!, I, K, L); Cianjur, Backer lowland alluvial forest, peat and mangrove swamp 23905, fr., 1-4-1918 (BO!). forest, beach forest, primary forest. 3. Licuala spinosa Wurmb. -Figure 4. Clustering. Notes: L. spectabilis is a synonym of L. spinosa Stem to 5 m or more tall, to c. 23 cm across. Leaves c. due to several characters such as: segments of frond, 15 to 17 in crown; sheaths disintegrating into coarse the number of inflorescence branching order % reticulate fibers with dark brown colours; inflorescence is branching to two orders, flowers hairy petioles to 2 m long, edge with black thorn, 10-20 mm and sessile. Miquel (1851), based his Licuala spectabilis near base, 5-10 mm toward apex, green to yellow on specimen collected by Junghuhn from Wijnkoops brownish; spines along whole length of petiole, triangu- Bay (close to Pelabuhan Ratu) in Banten, West Java % lar, patent to reflexed, largest near base; fronds peltate- where L. spinosa is also found. Miquel did not mention orbicular, dark green, c. 50-65 cm long, c. 80-150 cm where the specimens kept. I have tried to look for the wide, segment c. 14-19, all rather the same size; specimen in BO, but I did not find it. However, it was lateral segment 2-5 costulate, 41-51 x 4-9 cm; central likely that Miquel used the specimen kept in BO when segment slightly larger than rest, bipartitus he described this species. Unfortunately, the specimen (half to 5-7 cm toward leaf base), sometimes petiolulate, is now believed to be lost. c. 8-14 costulate, c. 52-65 x 20-23 cm. Inflorescences Specimen examined: WEST JAVA: P. Panaitan, erect to patent, longer than leaves, extend above the Waalkes 470, st., 10-9-1951 (BO!); Ujung Kulon, P. crown, with 7-9 partial inflorescences, c. 1.5-3 m long, Peucang, Dransfield 1478, fl., 18-4-1971 (BO!); Serpong, branching to two orders; prophyll tubular, c. 20 cm long Backer 17594, fr., s.d. (BO!); Kemayoran, Backer 427, or more, c. 1.5-2 cm wide, coriaceus, flattened, closely fl.+ fr., s.d. (BO!); Pelabuhan Ratu, Etti Siregar 01, fr., sheathing, apex with two keels, densely covered with 24-8-2002 (BO!), Dransfield 1098 (BO!, Ithaca); stellate caducous ferruginous hairs; peduncle c. Preanger, Koorders 34586β, fl., 14-5-1899 (BO!), 40-90 cm long, c. 4.5-8 across basally; peduncular bract Koorders 34589β, fl., 1-4-1899 (BO!); Cibunar, Dransfield present, sometimes lacking; rachis rigid, not sinuous; 1472, fl. 17-4-1971 (BO!); Dransfield 1407, fl., 13-4- rachis bracts similar to prophyll, c. 20-22 x 0.7-1.5 cm, 1971(BO!); Tanjung Suka, Van Steenis 994, fl., 3-4-1928 rachis bracts mouth splitting neatly in a few lobed; (BO!). CENTRAL JAVA: Karimun Jawa, Tg. Gelam, Karta rachillae 4-11 at one partial inflorescence, 207, fr., 23-11-1930(BO!), Karta 304, fr., 26-11-1930 unornamented, slightly close to or away from mouth of (BO!); Jepara, Ngarengan, Koorders 33618β, fl. 26-5- rachis bract, 10-25 cm long, c. 1.5-3 mm wide, covered 1899 (BO!), Koorders 36846β, fl., s.d., (BO!), Beümee with scattered simple brown hairs. Flowers solitary to 3598, fl., Dec. 1918 (BO!); Kedung Jati, Tempuran, in group of 2-4, sessile, densely arranged, cincinni Koorders 25443β, fr., 30-10-1896 (BO!); EAST JAVA: 5-10 per cm, maturing not simultaneously; bud c. Sukamade, Meru Betiri, Dransfield 3525, fl., 20-5-1973 4-5 x 2.5-3 mm; calyx cylindrical to cyathiform, c. (BO!, L). 3 x 2.5–3 mm, base thickened, apex trilobed, acumi- nate, lobed to about half of calyx length, covered with CONCLUSION translucent scattered patent hairs; corolla c. Three species of Licuala in Java are recognized. 3.5-4 x 3 mm, thick, densely covered in upper two third They are L. gracilis, L. pumila and L. spinosa. Two with simple translucent hairs, glabrescent toward base, previously known species have been placed into lobes acute c. 1.5 x 2 mm; staminal ring truncate c. synonymies; L. flabellum to L. gracilis, and L. 0.6-0.8 mm high, filament subulate, c.0.3 mm long, an- spectabilis to L. spinosa. thers c. 0.3 mm long; ovary glabrous, turbinate, apex truncate, 1.5 x 1 mm, style filiform, 1 mm long. Fruit ACKNOWLEDGEMENTS globose c. 6.5-8.5 x 6-8 mm, glabrous, smooth, imma- This paper forms part of the thesis for master ture fruit green and orange to red when mature. Seed degree at the Bogor Institute of Agriculture in 2004, globose, smooth, c. 4-6 mm across. with financial supports from DIKTI. I would like to thank Distribution: Andaman and Nicobar Islands, my supervisors Dr. Sri Sudarmiyati Tjitrosoedirdjo, Thailand, Vietnam, Malaysia, the Philippines, Sumatra, The genus Licuala Wurmb (Arecaceae) in Java 231 M.Sc. (IPB) and Dr. Johanis P. Mogea (LIPI) for their Dransfield, J. 1976. Palms in the every life of west Indonesia. Principes 20: 39-47. advice and guidance throughout the thesis. I also would Dransfield, J. & Moore, H.E. 1982. The Martian correlation two like to thank to the Herbarium Bogoriense for the loans editions of Martius’ Historia Naturalis Palmarum Compared. Kew Bull 37: 91-116. material. Greuter, W. et al. 2000. International Code of Botanical Nomen- clature (St Louis Code). Konigstein. Koeltz Scientific Books. Art. 13 REFERENCES Martius, C.F.P Von. 1838. Historia Naturalis Palmarum. Munich. Ardan, A.S. 2000. The Sumatran Licuala Revised. Thesis Post III: 119. Graduate. Bogor: IPB. Miquel, F.A.G. 1851. Plantae Junghuhnianae. Van Landbouw. 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